• mutants develop iris disease that is first noticeable at 6 months of age when eyes develop a pronounced peripupillary swelling accompanied by pronounced accumulation of clump cells on the iris surface

• double mutants develop iris disease that is first noticeable by 6 months of age, when all eyes show slight swelling of peripupillary tissue

• at 12, 14, and 18 months of age, increasing degree of iris atrophy is observed, that includes full-thickness iris holes, profound transillumination, pigment dispersion and frequent pigment accumulation on the lens and cornea, and changes to the dimensions of the anterior chamber

• however, mutants do not develop glaucomatus nerve damage that is seen in DBA/2J mice